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A new hadrosaurid dentary from the latestMaastrichtian of the Pyrenees (north Spain) and thehigh diversity of the duck-billed dinosaurs of the Ibero-Armorican Realm at the very end of the CretaceousPenélope Cruzado-Caballero a , José Ignacio Ruiz-Omeñaca a b c , Rodrigo Gaete d , VioletaRiera e , Oriol Oms e & José Ignacio Canudo aa Paleontología, Facultad de Ciencias, Grupo Aragosaurus-IUCA, Universidad de Zaragoza ,E-50009 , Zaragoza , Spainb Museo del Jurásico de Asturias (MUJA) , E-33328 , Colunga , Spainc Departamento de Geología , Universidad de Oviedo , c/ Jesús Arias de Velasco s/n,E-33005 , Oviedo , Spaind Museu de la Conca Dellà , C/ del museu, 4, E-25650 , Isona , Spaine Departament de Geologia , Universitat Autònoma de Barcelona, Cerdanyola del Vallès ,E-08193 , Barcelona , SpainPublished online: 02 Sep 2013.
To cite this article: Historical Biology (2013): A new hadrosaurid dentary from the latest Maastrichtian of the Pyrenees (northSpain) and the high diversity of the duck-billed dinosaurs of the Ibero-Armorican Realm at the very end of the Cretaceous,Historical Biology: An International Journal of Paleobiology
To link to this article: http://dx.doi.org/10.1080/08912963.2013.822867
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A new hadrosaurid dentary from the latest Maastrichtian of the Pyrenees (north Spain) and the highdiversity of the duck-billed dinosaurs of the Ibero-ArmoricanRealm at the very end of theCretaceous
Penelope Cruzado-Caballeroa1*, Jose Ignacio Ruiz-Omenacaabc2, Rodrigo Gaeted3, Violeta Rierae4, Oriol Omse5 and
Jose Ignacio Canudoa6
aPaleontologıa, Facultad de Ciencias, Grupo Aragosaurus-IUCA, Universidad de Zaragoza, E-50009 Zaragoza, Spain; bMuseo delJurasico de Asturias (MUJA), E-33328Colunga, Spain; cDepartamento deGeologıa, Universidad deOviedo, c/ Jesus Arias de Velasco s/n,E-33005 Oviedo, Spain; dMuseu de la Conca Della, C/ del museu, 4, E-25650 Isona, Spain; eDepartament de Geologia, UniversitatAutonoma de Barcelona, Cerdanyola del Valles, E-08193 Barcelona, Spain
(Received 6 May 2013; final version received 3 July 2013)
In the latest Maastrichtian, the European hadrosauroid fauna was more diverse than those of North America and Asia. TheEuropean record of hadrosaurid dentaries is an example of this diversity, and most of the sites with mandibular remains arelocated in the Ibero-Armorican Realm. Within the Iberian Peninsula, most of the remains are located in the Tremp Basin(South Central Pyrenees). Two of the three valid hadrosaurid taxa defined in this basin are from the Blasi sites (Aren, Huescaprovince): Arenysaurus ardevoli (Blasi-3) and Blasisaurus canudoi (Blasi-1). A new locality in Blasi (Blasi 3.4) hasprovided a new dentary from an indeterminate euhadrosaurid. This dentary presents some characters intermediate betweenArenysaurus and Blasisaurus, some characters similar to Pararhabdodon isonensis (from the nearby province of Lleida),and some characters of its own. Nevertheless, due to its fragmentary character, without dentition or its edentulous anteriorpart, it cannot be determined above the level of Euhadrosauria. It thus represents a fourth Iberian euhadrosaurian taxon in theIbero-Armorican Realm, different from Arenysaurus, Blasisaurus and Pararhabdodon, increasing the diversity ofhadrosauroids in this realm at the very end of the Cretaceous.
Keywords: Hadrosauridae; Late Cretaceous; dentary; paleobiodiversity; Iberian Peninsula
Introduction
The paleobiodiversity of the European duck-billed dinosaurs
of the Maastrichtian distinguishes them from those from
North America and Asia (Prieto-Marquez et al. 2007; Dalla
Vecchia 2009; Pereda-Suberbiola, Canudo, Company, et al.
2009; Pereda-Suberbiola, Canudo, Cruzado-Caballero, et al.
2009; Prieto-Marquez and Wagner 2009; Cruzado-
Caballero, Pereda-Suberbiola and Ruiz-Omenaca 2010;
Cruzado-Caballero, Ruiz-Omenaca and Canudo 2010). This
mixedEuropean faunawas due to the connection ofAsia and
Europe by means of ‘semi-permeable’ barriers that were
emerged or submerged depending on changes in the climate
and sea level (Prieto-Marquez and Wagner 2009). A series
of geo-dispersal events is likely to have occurred. The first
event probably took place during the Santonian–Campanian
of Asia and explains the presence of the non-hadrosaurid
hadrosauroid Tethyshadros insularis Dalla Vecchia, 2009
and the basal hadrosaurid Telmatosaurus transsylvanicus
(Nopcsa, 1903) (Cruzado-Caballero, Pereda-Suberbiola
et al. 2010). Afterwards, no later than the middle to late
Campanian, another dispersal event of hadrosaurines and
lambeosaurines is likely to have taken place. The periods of
isolation during high-sea-level times may have facilitated
the survival of non-hadrosaurid hadrosauroids and basal
hadrosaurids (i.e. Tethyshadros and Telmatosaurus) in the
Maastrichtian of Europe (Pereda-Suberbiola, Canudo,
Company, et al. 2009). These taxa took refuge in the
European islands and underwent varying degrees of
reduction in size probably due to insular dwarfism (Dalla
Vecchia 2009; Prieto-Marquez and Wagner 2009; Benton
et al. 2010; Weishampel et al. 2010). As a result of the
different geo-dispersal events and the periods of isolation,
the European faunawas the only onewhere non-hadrosaurid
hadrosauroids, non-euhadrosaurian hadrosaurids, hadro-
saurines and lambeosaurines co-existed in theMaastrichtian.
In other continents (Asia, North America and South
America) the more basal forms became extinct before the
Campanian.
The European record is based on many fragmentary
specimens with a good representation of dentaries (from
France, Italy, Romania, Spain and The Netherlands;
Figure 1). Most of the sites with remains of dentaries are
located in the Pyrenees (south France and north Spain) and
are Maastrichtian in age (Table 1). The French record is
very fragmentary, and most of it probably belongs to
juvenile specimens (Laurent et al. 2002; Laurent 2003),
while the Spanish record presents at least 13 specimens of
adult and juvenile individuals. Three of these dentaries
come from the Blasi sites in Aren (Huesca province;
Cruzado-Caballero, Ruiz-Omenaca, Gaete et al. 2010;
Figure 2). The Blasi sites are the richest in cranial and
q 2013 Taylor & Francis
*Corresponding author. Email: penelope@unizar.es
Historical Biology, 2013
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postcranial hadrosaurid remains described in the latest
Maastrichtian of Spain (Lopez-Martınez et al. 2001;
Cruzado-Caballero et al. 2005; Pereda-Suberbiola, Canudo,
Cruzado-Caballero, et al. 2009; Cruzado-Caballero, Pereda-
Suberbiola and Ruiz-Omenaca 2010; Cruzado-Caballero,
Ruiz-Omenaca and Canudo 2010). The aim of this paper is
to describe a new dentary from the new locality Blasi 3.4
and to try to assess the diversity of the European
hadrosauroids of the Latest Cretaceous on the basis of the
preserved dentaries.
Geological and chronological framework
TheTrempGroup (see references inRosell et al. 2001) is part
of the south-Pyrenean foreland basin infill that took place
during the Upper Cretaceous and Paleogene. This formation
is the result of sediment deposition in continental settings
(fluvial environments) and transitional marine-continental
settings (mainly lagoonal environments). Such stratigraphic
units change laterally to each other aswell aswith themarine
Aren Formation (coastal environments). The age of the
Tremp Group (Oms et al. 2007; Riera et al. 2009) ranges
from Maastrichtian to Paleocene, and in the area of the
studied material it has been dated by means of magnetos-
tratigraphy (Oms and Canudo 2004; Pereda-Suberbiola,
Canudo, Cruzado-Caballero, et al. 2009; Vila et al. 2012).
Thus all the findings reported here can be integrated within a
robust chronostratigraphic framework to show that the levels
with hadrosauroid mandibles are found in a stratigraphic
position close to one another (see Figure 3) around the
boundary between chrons C29r and C30n (i.e. ca. 66.2Ma).
Institutional abbreviations
IPS, Institut Catala de Paleontologia (formerly Institut de
Paleontologia DrMiquel Crusafont), Sabadell, Spain;MDE,
Musee des dinosaures, Esperaza, France; MGUV,Museo de
Geologıa,Universitat deValencia, Valencia, Spain;MNHN,
Museum national d’Histoire naturelle, Paris, France; MPV,
Museo Municipal de Ciencias Naturales de Valencia,
Valencia, Spain; MPZ, Museo Paleontologico de la
Universidad de Zaragoza, Zaragoza, Spain.
The Ibero-Armorican record of dentaries from the
Maastrichtian
The Ibero-Armorican record consists of 19 dentaries,
6 from France and 13 from Spain (Figure 4).
The French record
Of the six French dentaries described, three are well
preserved, whereas the other three are very fragmentary.
The three best-preserved dentaries comprise two from the
site of Cassagnau-2 (Marignac-Laspeyres, Department of
Haute-Garonne; MDE-Cas2-02, 248) and one from the site
of Le Bexen (Fontjoncouse, Department of Aude; MDE-
Fo1-10). According to Laurent (2003), all of them belong
to juvenile individuals.
An incomplete right dentary (MDE-Cas2-02, Hadro-
sauridae indet.) and a complete left dentary (MDE-Cas2-
248, Euhadrosauria indet.) were found in the site of
Cassagnau-2. This site is dated as upper Maastrichtian
[AuzasMarls Formation;Laurent et al. 2002;Laurent 2003].
Both dentaries are gracile and anteroposteriorly short;
MDE-Cas2-248 has a length of 160mm (Figure 4(B)).
The coronoid process is slightly projected anteriorly, and the
posterodorsal edge is higher than the anterodorsal edge. In
lateral view, several nutritional foramina are observed,
and in MDE-Cas2-248 there is a large nutritional
foramen near the symphyseal region. The dental battery of
Figure 1. Simplified paleogeographical map of Europe during the Late Cretaceous (Maastrichtian), redrawn from Pereda-Suberbiola(2009, fig. 2), with the distribution of the known European hadrosauroid dentaries. A, Apennine platform; AA, Australpine island; AD,Adriatic-Dinaric island; AI, Alboran islands; Ap, Apulian platform; AS, Anglo-Scottish island; C, Crimean island; FS, Fennoscandianshield; I, Irish island; IA, Ibero-Armorican island; RB, Rhenish-Bohemian island; Rh, Rhodope; T, Transylvanian basin; US, Ukrainianshield. Plot marks: forest, exposed land; dotted, epicontinental seas; dark grey, oceanic basins; striped, unknown; black, volcanics.
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Table
1.
LatestCretaceoushadrosauroid
dentary
record
from
Europe.
Site/country
Form
ation
Age
Taxon
Rem
ains
Reference
Ankerpoort-’tNekam
iQuarry,province
of
Lim
burg,TheNetherlands
MaastrichtForm
ation,
possibly
Nekum
Mem
ber
Upper
Maastrichtian
Euhadrosauriaindet.
NHMM
198027Fragmentary
rightdentary
Buffetautet
al.(1985),
Dalla
Vecchia
(2006)
Sanpetru,Hateg
Basin,
Transsylvania,Romania
Sanpetru
Fromation
Maastrichtian
Telmatosaurus
transsylvanicus
BMNH
3386Holotype:
askull
Nopcsa(1903),Weisham
pel
etal.(1993),Dalla
Vecchia
(2006)
LeJadet,Departm
entof
Haute-G
aronne,France
Calcarenites
duJadet
Form
ation(A
usseing
Group)
Upper
Maastrichtian
Hadrosauridae
indet.
MNHNP-sma1Fragmentary
leftdentary
ParisandTaquet
(1973),
Laurent(2003)
Cassagnau
2,Departm
ent
ofHaute-G
aronne,France
Marnes
d’A
uzas
Form
ation
Upper
Maastrichtian
(upper
Cam
panian/upper
Maastrichtian
inLopez-
Martınez
etal.2001)
Hadrosauridae
indet.
Euhadrosauriaindet.
MDE-Cas2-02Rigth
dentary;
MDE-Cas2-248Leftdentary
Laurent(2003)
LeBexen,Departm
ent
ofAude,France
Marnes
Rouges
de
RoquelongueForm
ation
Upper
Maastrichtian
(middle
Maastrichtian
inLopez-M
artınez
etal.2001)
Hadrosauridae
indet.
MDE-Fo1-01Fragmentary
rightdentary;MDE-Fo1-03
Fragmentary
leftdentary;
MDE-Fo-10Fragmentary
rightdentary
Laurentet
al.(1997),
Laurent(2003)
Villaggio
del
Pescatore,
Trieste,Italy
LiburnianForm
ation
Upper
Cam
panian–
Paleocene
Tethyshadrosinsularis
SC57021Holotype:
acomplete
andarticulated
skeleton
Dalla
Vecchia
(2009)
Fontllonga,Province
of
Lleida,Spain
Conques?Form
ation
(TrempGroup)
Upper
Maastrichtian
Hadrosauroidea
indet.
IPS36338Fragmentary
leftdentary
Casanovas-Cladellas
etal.(1999b),Pereda-Suberbiola,
Canudo,Cruzado-Caballero
etal.
(2009)
Les
Llaus,Province
ofLleida,Spain
Conques
Form
ation
(TrempGroup)
Upper
Maastrichtian
Koutalisaurus
kohleroruma
IPS-SRA-27Holotype:
arigth
dentary
Casanovas-Cladellas
etal.(1999a),Prieto-M
arquez
andWagner
(2009)
Basturs
Poble,Province
ofLleida,Spain
Conques
Form
ation
(TrempGroup)
Upper
Maastrichtian
Hadrosauridae
indet
Atleastthreerigth
dentaries
Prieto-M
arquez
etal.(2007)
LaSolana,Province
ofValencia,Spain
Villalbadela
Sierra
Form
ation
Upper
Maastrichtian
Hadrosauridae
indet.
MGUV
2200Leftdentary;
MPV
181Rightdentary;
MPV
182Fragmentary
rightdentary;
MPZ99/665Leftdentary.
CompanyRodrıguez
(2004),
Pereda-Suberbiola,Canudo,
Cruzado-Caballero
etal.(2009)
Blasi1,Province
of
Huesca,Spain
ArenForm
ation
Upper
Maastrichtian
Blasisauruscanudoi
Lopez-M
artınez
etal.(2001),
Cruzado-Caballero,Pereda-
Suberbiola
and
Ruiz-O
menaca
(2010)
Blasi3,Province
of
Huesca,Spain
LaPosa
Form
ation
(TrempGroup)
Upper
Maastrichtian
Arenysaurusardevoli
MPZ2008/258;leftdentary
Pereda-Suberbiola,Canudo,
Cruzado-Caballero
etal.(2009)
Blasi3.4,Province
of
Huesca,Spain
LaPosa
Form
ation
(TrempGroup)
Upper
Maastrichtian
Euhadrosauriaindet.
MPZ2010/952;leftdentary
Thispaper
Barranco
Serraduy1,
Province
ofHuesca,Spain
LaPosa
Form
ation
(TrempGroup)
Upper
Maastrichtian
Hadrosauridae
indet.
BS1-13;Fragmentary
leftdentary
Puertolas-Pascual
etal.(2012)
Amor2,Province
of
Huesca,Spain
Conques
Form
ation
(TrempGroup)
Upper
Maastrichtian
Hadrosauridae
indet.
MPZ2013/72;rightdentary
Cruzado-Caballero
etal.(2012)
aThisdentary
was
designated
theholotypeofKoutalisaurusbyPrieto-M
arquez
etal.(2006);now
ajuniorsubjectivesynonym
ofPararhabdodon(Prieto-M
arquez
andWagner,2009).
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MDE-Cas2-248 is made up of 21 tooth positions with two
preserved replacement teeth, and MDE-Cas2-02 has 21
tooth positions but no teeth are preserved. The teeth of
MDE-Cas2-248 have a single prominent median carina that
is straight and slightly caudally placed, and they do not have
secondary ridges. Laurent et al. (2002) proposed that MDE-
Cas2-248 is very similar to Blasisaurus (as Euhadrosauria
indet. in Lopez-Martınez et al. 2001, fig. 5(C),(D)), sharing a
gracile form, the absence of secondary ridges and a coronoid
process that is slightly projected anteriorly.
An incomplete right dentary (MDE-Fo1-10, Hadrosau-
ridae indet.; Figure 4(A)) from the site of Le Bexen (Marnes
Rouges de Roquelongue Formation; upper Maastrichtian)
was described byLaurent et al. (1997). It is the largest French
dentary (it is 210mm long) but it does not preserve the
anterior and posterior ends and the coronoid process. The
Meckelian canal is large, and the suprameckelian foramen is
oblique, at 458 to the alveolar medial surface. This dentary
preserves 24 tooth positions without teeth. The other two
dentaries described from Le Bexen (MDE-Fo1-01, MDE-
Fo1-03) are very fragmentary (see Laurent et al. 1997). One
additional dentary from the French record (MNHNP-sma 1
from the locality of Le Jadet) is too fragmentary for
comparisons to be drawn. This is late Maastrichtian in age
and is assigned to Hadrosauridae indet. (See Table 1).
The Spanish record
In the Iberian Peninsula, the paleobiodiversity of
hadrosaurids can be said to be high on the basis of the
preserved dentaries. All the localities but one (La Solana in
Figure 2. Setting of the studied site and sections with a general geographic location (A) and indication of the sketched geological map ofthe Pyrenees (B). The latter also indicates the map with the precise location of the sections (C) that can be found in Figure 3.
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the South-western Iberian Range; Company Rodrıguez
2004; Pereda-Suberbiola, Canudo, Company, et al. 2009)
are in the South Central Pyrenees. According to the
dentary remains, the Spanish Maastrichtian record consists
of at least five different taxa: (1) an indeterminate non-
hadrosaurid hadrosauroid from Fontllonga site (IPS
36338, Figure 4(C); Camarasa municipality, Lleida
province), (2) an indeterminate hadrosaurid from La
Solana site (MGUV 2200 and MPV 181, Figure 4(E),(F);
Tous municipality, Valencia province), (3) Koutalisaurus
Prieto-Marquez, Gaete, Rivas, Galobart and Boada, 2006
from Les Llaus site (IPS SRA 27, Figure 4(D); Abella de la
Conca municipality, Lleida province; junior synonym of
Pararhabdodon Casanovas-Cladellas, Santafe-Llopis and
Isidro-Llorens 1993 after Prieto-Marquez and Wagner
(2009)), (4) Arenysaurus Pereda-Suberbiola, Canudo,
Cruzado-Caballero, Barco, Lopez-Martınez, Oms and
Ruiz-Omenaca, 2009 from Blasi 3 site (MPZ2008/258;
Figure 4(H); Aren municipality, Huesca province) and (5)
Blasisaurus Cruzado-Caballero, Pereda-Suberbiola and
Ruiz-Omenaca, 2010 from Blasi 1 site (MPZ99/664,
Figure 4(G); Aren municipality) (see Pereda-Suberbiola,
Figure 3. (Colour online) Location of the studied mandible (Blasi 3.4 site), in the west area of Blasi (section 5) and correlation withsections 4 to 1 (Blasi-Aren and Isona areas). See location of the sections in Figure 2(C). Lithostratigraphic correlations combined withmagnetostratigraphic data from section 1, and Isona and Vallcebre, clearly indicate the correlation of Blasi 3.4 with the upper part ofchron C30n (late Maastrichtian).
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Canudo, Company, et al. 2009; Pereda-Suberbiola,
Canudo, Cruzado-Caballero, et al. 2009; Prieto-Marquez
and Wagner 2009; Cruzado-Caballero, Pereda-Suberbiola
et al. 2010 and references therein).
Several additional dentaries have been discovered, all
of them currently indeterminate hadrosaurids. These are
from the sites of Basturs Poble (Isona i Conca Della
municipality, Lleida province; Prieto-Marquez et al.
2007), Amor-2 (Beranuy municipality, Huesca province;
Cruzado-Caballero et al. 2012) and Barranco Serraduy-1
(Isabena municipality, Huesca province; Puertolas-Pas-
cual et al. 2012).
IPS 36338 (Figure 4(C)) is a fragmentary left dentary
from Fontllonga site (upper Maastrichtian; Conques
Formation, Tremp Group). It is robust, with a very strong
sub-vertical coronoid process, whose posterodorsal side is
large and protruding. In posterior view, the supramec-
kelian foramen is oblique. The dental battery does not
extend posteriorly to the coronoid process. In medial view,
it is possible to make out 30–31 tooth positions, and each
Figure 4. Silhouette of the Late Cretaceous Ibero-Armorican island (see Figure 1) with the location of the best-preserved hadrosauroiddentaries; star: Le Bexen (Aude department) and Cassagnau 2 (Haute-Garonne department); circle: Blasi sites (Huesca province);triangle: Fontllonga and Les Llaus (Lleida province); square: La Solana (Valencia province). A–H: dentaries in medial view; all are leftdentaries except A, D and F, which are right dentaries that have been turned around for better comparison. A: MDE-Fo1-10,Hadrosauridae indet. from Le Bexen; B: MDE-Cas2-248, Euhadrosauria indet. from Cassagnau 2; C: IPS-36338, Hadrosauroidea indet.from Fontllonga; D: IPS-SRA-27, Pararhabdodon isonensis from Les Llaus; E: MGUV-2200, Hadrosauridae indet. from La Solana; F:MPV-181, Hadrosauridae indet. from La Solana; G: MPZ99/664 Blasisaurus canudoi from Blasi 1; H: MPZ2008/258, Arenysaurusardevoli from Blasi 3. C, E, F taken from Pereda-Suberbiola, Canudo, Company et al. (2009, fig. 1(E)–(G)); D taken from Prieto-Marquez et al. (2006, fig. 7(A)); G taken from Cruzado-Caballero, Pereda-Suberbiola and Ruiz-Omenaca (2010, fig. 3.1); H taken fromPereda-Suberbiola, Canudo, Cruzado-Caballero et al. (2009, fig. 4(A)). Scale bar: 5 cm.
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position has two functional teeth and two replacement
teeth. The ventral deflection of the dentary (sensu Horner
et al. 2004) is located at about the 12th most anterior tooth
position. The teeth present a faint secondary ridge that,
according to Casanovas-Cladellas et al. (1999b), could be
bifid in the basal part of the crown. Also, these teeth have
neither papillae nor denticles.
MGUV 2200 and MPV 181 (Figure 4(E),(F)) are two
fragmentary dentaries from juvenile and adult individuals,
respectively. These dentaries were found in the site of La
Solana (upper Maastrichtian; Villalba de la Sierra
Formation). In both dentaries the dental battery terminates
posterior to the apex of the coronoid process. Also in both
dentaries the deflection is located at about the 11–12th
most anterior tooth position. The coronoid process is
slender, sub-vertical, and in posterior view an oval and
oblique suprameckelian foramen can be seen.
MGUV 2200 has 29 tooth positions with some teeth
preserved. The teeth have a faint secondary ridge and bear
small marginal denticles not supported by ridges. MPV 181
has 36–37 tooth positions and preserves some shifted teeth.
IPSSRA27 is the only knowndentary currently assigned
toPararhabdodon. It comes from the site ofLesLlaus (upper
Maastrichtian; Conques Formation, Tremp Group), 750m
west of the site of Sant Roma d’Abella (Isona and Conca
Della municipality), the type locality of Pararhabdodon
(Prieto-Marquez et al. 2006). It is a complete right dentary
with a contact surface for the predentary that is extremely
long and strongly deflected (Figure 4(D)). The coronoid
process is broken, so it does not fitwith themandibular ramus
and its current backward inclination is an artefact;
nevertheless it is complete and has a height of 104mm
from its base to its apex. The dental battery has a length of
243mm and shows 35 tooth positions but does not preserve
any teeth (Casanovas-Cladellas et al. 1999a; Prieto-Marquez
et al. 2006; Prieto-Marquez and Wagner 2009). In occlusal
view, the dental battery is straight but it turns slightly inwards
in the last positions, i.e. those adjacent to the coronoid
process. The deflection point of the dentary is located
between the 10–11th tooth positions.
Two lambeosaurines have beendescribed from the upper
Maastrichtian Blasi sites inAren:Blasisaurus from the Blasi
1 site (ArenFormation) andArenysaurus from theBlasi 3 site
(La Posa Formation, Tremp Group). Both specimens
preserve the left dentary. The dentary of Blasisaurus
(MPZ99/664, Figure 4(G)) is slightly less robust than that of
Arenysaurus (MPZ2008/258, Figure 4(H)). In both dentaries
the coronoid process is anteriorly projecting. In Blasisaurus
the dorsal side of the coronoid process is convex, and in
Arenysaurus it is tip-like with the posterodorsal border
higher than the anterodorsal border. Also, in these dentaries
the anteriormost portion is modestly deflected ventrally. The
dental battery of Blasisaurus has a length of 170mm and 35
tooth positions, and that of Arenysaurus has a length of
275mm and 37 tooth positions. In the Arenysaurus dentary
the starting point of the deflection is located at the 9–10th
tooth position, whereas in the Blasisaurus dentary it starts at
the 10–11th tooth position. The teeth of both taxa have a
slightly sinuous primary ridge and do not have papillae. The
Arenysaurus teeth have a posterior secondary ridge that is
absent in Blasisarus.
In the bonebed of Basturs Poble (upper Maastrichtian,
Conques Formation, Tremp Group), there are disarticu-
lated cranial and postcranial remains of at least five
hadrosaurid individuals. These have been cited in an
abstract but have not been described in detail or figured
(Prieto-Marquez et al. 2007). The dentaries have a rostral
end that is elongated medially and nearly as long as that of
Pararhabdodon. Some teeth have secondary ridges. There
are at least three dentaries (unpublished data) but
unfortunately there are no adult dentaries in the sample;
Prieto-Marquez et al. (2007) say that ‘these specimens
may represent Pararhabdodon; however, the discovery of
an adult dentary is required to test the hypothesis’.
MPZ2013/72 is a fragmentary right dentary probably
from a juvenile individual from the site of Amor-2 (upper
Maastrichtian, Conques Formation, Tremp Group) near
the village of Beranuy (figured erroneously as AM3-15
(from the Amor-3 site) by Cruzado-Caballero et al. (2012,
fig. 2(A)). This dentary only preserves eight tooth
positions without teeth and the base of the coronoid
process. The Meckelian canal has an ellipsoidal form and
is oblique to the alveolar medial surface.
Another dentary from the Beranuy area (Barranco
Serraduy-1 site, upper Maastrichtian, La Posa Formation,
TrempGroup) has been cited in a faunal list for the Beranuy/
Serraduy sites (Puertolas-Pascual et al. 2012, table 1).
BS1-13 is a left dentary, but it remains unprepared so it is not
possible to ascertain its characteristics at present.
Recently, a new site has been found in the Blasi area
(Aren; Blasi 3.4), and from its surface a new hadrosaurid
dentary has been collected (MPZ2010/952; Cruzado-
Caballero, Ruiz-Omenaca, Gaete et al. 2010, fig. 3). This
is described in this paper.
Systematic paleontology
Superorder Dinosauria Owen, 1842
Order Ornithischia Seeley, 1887
Infraorder Ornithopoda Marsh, 1881
Family Hadrosauridae Cope, 1870
Euhadrosauria Weishampel et al., 1993
Euhadrosauria indet
(Figure 5)
Material. A left dentary (MPZ2010/952) from the Blasi
3.4 site (Aren, Huesca province), upper Maastrichtian, La
Posa Formation (Tremp Group).
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Description
MPZ2010/952 is an incomplete left dentary (Figure 5(A)–
(F)) that does not preserve the symphyseal zone. It has
several diagnostic characters ofHadrosauridae sensuHorner
et al. (2004), such as a distal extension of the dentary tooth
row caudal to the apex of the coronoid process, the dental
battery inclined downwards in its anterior part and the
caudalmost end of the dentary well behind the coronoid
process, which is inclined slightly forwards (Figure 5(A)).
MPZ2010/952 has parallel dorsal and ventral borders.
In medial view the angular facet is broken, and the
suprameckelian foramen forms an angle of about 308 withthe alveolar medial surface, similar to IPS 36338 and
MGUV 2200, and unlike MDE-Fo1-10 and Telmatosaurus
with 908 (Figure 5(C), Casanovas-Cladellas et al. 1999b;
Laurent et al. 2002). Through this foramen passes the
external mandibular artery that covers the whole lateral side
of the dentary and ends in the large foramen of the
symphyseal zone. From this artery at least five small blood
vessels issue that end in the nutritional foramina that
perforate the lateral side (Figure 5(B),(D),(E)). The external
mandibular artery runs parallel to the internal mandibular
artery, which covers theMeckelian canal. In dorsal view, the
longitudinal axis of the dentary is parallel to the lateral side
and is slightly arched lingually (Figure 5(F)).
The dental battery does not have any teeth; it is
complete, with an anteroposterior length of 225mm and
36 dental positions (Figure 5(A)). The dental battery is
shorter than that of Arenysaurus (275mm) and longer than
that of Blasisaurus (170mm). The number of tooth
positions is similar to MPV 181 with 36–37 positions, less
than Arenysaurus with 37, and greater than Blasisaurus
and Pararhabdodon with 35, IPS 36338 with less than
34 and MGUV 2200 with 29 (Company Rodrıguez 2004;
Pereda-Suberbiola, Canudo, Company, et al. 2009; Pereda-
Suberbiola, Canudo, Cruzado-Caballero, et al. 2009;
Cruzado-Caballero, Pereda-Suberbiola and Ruiz-Ome-
naca 2010). In dorsal view, the dental battery is straight
with its posteriormost part deflected medially, similar to
Pararhabdodon. The tooth positions have a width of 6mm,
and are straight except in the proximal half, where they
incline anteriorly. These positions end posterior to the apex
Figure 5. Euhadrosauria indet. from Blasi 3.4. MPZ2010/952 left dentary in medial (A), lateral (B, D, E), posterior (C) and dorsal (F)views. B, D and E show different details of the nutritional foramina, with the external mandibular artery covered (B, D) and exposed (E).White arrows in D indicate the blood vessels that end in the nutritional foramina.
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of the coronoid process as in Pararhabdodon. Although
the ventral border is broken, it has been possible to
estimate a midpoint depth of 65.36mm in medial view
(Figure 6). Due to the breakage of the ventral border,
moreover, it is possible to see the internal tissue fibres,
which are deflected ventrally in the 13–14th tooth
positions of the dental battery. This arrangement of fibres
marks the start of the anterior ventral deflection of the
dentary.
The coronoid process is high (112mm in length),
slender, with a flat inner side as is usual in hadrosaurids; it
is more anteriorly inclined than Arenysaurus and
Blasisaurus (but not as strongly projected as shown in
Figure 5(A) due to preservation problems) and slightly
curved inwards (Figure 5(C)). Anterodorsally the coronoid
process is broken and it is not possible to know whether it
is expanded anteriorly. Like Telmatosaurus, the ante-
rodorsal border seems to be higher than the posterodorsal
border, unlike Arenysaurus, Charonosaurus jiayinensis
Godefroit, Zan and Jin, 2000, Amurosaurus riabinini
Bolotsky and Kurzanov, 1991 and MGUV 2200, where the
posterodorsal side is higher than the anterodorsal side,
and unlike Blasisaurus and Sahaliyania elunchunorum
Godefroit, Hai, Yu and Lauters, 2008, where both borders
are equal in height (Weishampel et al. 1993; Godefroit
et al. 2001, 2004, 2008; Company Rodrıguez 2004). The
adductor fossa is narrow.
Discussion
As we consider MPZ2010/952 to be the dentary of an adult
individual, here it will only be compared with other adult
dentaries from the Ibero-Armorican record (i.e. Spanish
record, as all the well-preserved French dentaries are from
juvenile individuals; see Laurent et al. 2002).
MPZ2010/952 has a relatively high number of dental
positions in the dental battery (36 in 225mm). In this
respect it falls between Blasisaurus (35 in 170mm) and
Pararhabdodon (35 in 263mm) and Arenysaurus (37 in
275mm). All these three dentaries are considered to be
non-juvenile lambeosaurine individuals. Although
MPZ2010/952 has no diagnostic character that identifies
it as a basal hadrosaurid, a lambeosaurine or a
hadrosaurine, it has a greater number of dental positions
than basal hadrosaurids [30 or less according to Godefroit
et al. (2008, char. 41) and Prieto-Marquez (2010, char. 1);
32 or less according to Horner et al. (2004, char. 1), Suzuki
et al. (2004, char. 46), Evans and Reisz (2007, char. 60)
and Sues and Averianov (2009, char. 92)]. Moreover, its
36 dental positions are within the range of the adult
individuals of hadrosaurines and lambeosaurines ((more
than 32 according to Horner et al. (2004, char. 1), Suzuki
et al. (2004, char. 46), Evans and Reisz (2007, char. 60)
and Sues and Averianov (2009, char. 92)). On the other
hand, the dentary tooth density (number of tooth positions
per cm; Prieto-Marquez 2008, char. 2) of MPZ2010/952 is
1.68, which according to this author lies within the range
of nearly all hadrosaurine and all lambeosaurine
hadrosaurids (except Charonosaurus), which have a
value .1.1 for this index. Moreover, it falls within the
range of Iberian lambeosaurines, which extends from 1.34
in Arenysaurus to 2.06 in Blasisaurus (Table 2). In
summary, the number of dental positions and their density
indicate that MPZ2010/952 is the dentary of an adult
hadrosaurid and probably a derived one.
According to Godefroit et al. (2008), although it is
variable, a ventral deflection of the rostral part of
the dentary is a usual character in lambeosaurines. The
position of the internal tissue fibres in the first third of the
dental battery of MPZ2010/952, together with the number
and the density of dentary teeth, indicates that the dentary
belongs to the lambeosaurine clade or at least affirms that
this dentary is euhadrosaurian (sensu Weishampel et al.
1993).
In order to ascertain whether or not the new dentary
from Blasi 3.4 belongs to one of the euhadrosaurian
species already known from the Ibero-Armorican Realm,
Figure 6. Schematic drawings of left dentary in medial (A) and lateral (B) views, showing the measurements taken for Table 2.H, heightof the dental battery at its midpoint, taken from the base of the dental battery to the uppermost dorsal border of the dentary, withoutmeasuring the functional crowns; L, length of the dental battery; h, height of coronoid process, measured in lateral view from its base to itsdorsalmost expansion. The arrow marks the relative position of the inflexion point where the ventral margin of the dentary originates (R).The dentary represented, as a model, in this figure is that of A. ardevoli, drawn by the authors (see Figure 4(H) for a picture). Scratchedareas represent matrix.
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several measurements have been taken in the dentaries of
these species (Table 2, see methods in Figure 6).
It is observed that the tooth density decreases when the
length of the dental battery increases, so this is not
discriminant as a specific character. Several features and
ratios distinguish MPZ2010/952 from the morphology of
both Arenysaurus and Blasisaurus dentaries, making it
more similar to the dentary of Pararhabdodon: (1) the
dental battery exceeds the posterior edge of the coronoid
process (in Arenysaurus it does not reach the posterior
edge, and in Blasisaurus it finishes just at the posterior
edge of the coronoid process), (2) in dorsal view the dental
battery is straight, with a small posterior zone that is
inclined medially (in Arenysaurus it is laterally concave
and in Blasisaurus it is straight but with a small anterior
zone laterally inclined), (3) the ratio of the location of the
origin of the ventral deflection of the dentary (Prieto-
Marquez 2010, char. 37) is 0.58 (Pararhabdodon 0.54,
Arenysaurus 0.65 and Blasisaurus 0.66), (4) the midpoint
height/total length ratio of the dental battery is estimated at
0.29 (Arenysaurus 0.26, Pararhabdodon 0.31 and
Blasisaurus 0.34).
MPZ2010/952 also differs from Arenysaurus and
Blasisaurus in having a coronoid process that is more
anteriorly sloping (in Pararhabdodon the orientation of the
coronoid process is an artefact due to breakage and plaster
reconstruction).
One of the ratios that best distinguishes MPZ2010/952
from Pararhabdodon is that between the height of the
coronoid process and the height of the dental battery at its
midpoint; in Pararhabdodon and Arenysaurus this ratio is
,1.4, and in Blasisaurus it is .2, whereas in
MPZ2010/952 its value is intermediate (1.71) (Table 2).
To a lesser extent, the ratio between the height of the
coronoid process and the length of the dental battery also
distinguishes the taxa; in Arenysaurus and Pararhabdodon
this ratio is,0.43; and in Blasisaurus it is.0.85, while in
MPZ2010/952 it is 0.50 (Table 2). In the light of these
ratios, together with the features and measures already
discussed, it seems most plausible that MPZ2010/952
represents a fourth euhadrosaurian taxon in the Ibero-
Armorican Realm, in addition to Pararhabdodon,
Arenysaurus and Blasisaurus.
According to Prieto-Marquez (2010, char. 36), on the
other hand, the ventral deflection of MPZ2010/952 lies
within the range of lambeosaurine values (0.65 or less),
with the exception of the hadrosaurines Brachylopho-
saurus canadensis Sternberg, 1953 and Gryposaurus
latidens Horner, 1992. This being so, it is more probable
that MPZ2010/952 belongs to a lambeosaurine than to a
hadrosaurine. It should be noted, however, that an
indeterminate hadrosaurine cranial bone has been
described in the nearby Blasi 5 site [Conques Formation;
a fragmentary right jugal (MPZ2007/1885) described by
Cruzado-Caballero, Ruiz-Omenaca and Canudo 2010], so
a hadrosaurinae origin cannot be ruled out.
Given the fragmentary character of MPZ2010/952, it is
not possible to assign it either to Lambeosaurinae or
Hadrosaurinae, but it can be recognised as an indetermi-
nate euhadrosaurian. It represents the fifth euhadrosaurian
taxon in the South Central Pyrenees and in the Ibero-
Armorican Realm.
Conclusions
The Ibero-Armorican record of hadrosauroid dentaries
from the Maastrichtian is the most complete and most
diverse in Europe, with at least five different taxa, all from
the Iberian Peninsula. This record presents dentaries of
both basal and derived taxa, and different ontogenetic
states. The high heterogeneity of the taxa recorded in the
Latest Cretaceous of Europe is thus the most complex in
Laurasia.
A new locality in the Tremp Basin of the South Central
Pyrenees (Blasi 3.4; Aren municipality, Huesca province,
Spain) has provided a new dentary that presents a mosaic
of characters that allow it to be assigned to indeterminate
euhadrosaurians. This dentary represents a different taxon
from Arenysaurus, Blasisaurus and Pararhabdodon, the
three euhadrosaurians already known in this basin.
This new dentary adds to the dentary record from
Europe and increases the great diversity of hadrosauroids,
Table 2. Measurements and indexes in selected dentaries of Late Cretaceous Ibero-Armorican euhadrosaurians, ordered by the length ofthe dental battery.
Material Taxon L H h n Td (n/L) H/L h/H h/L R
MPZ99/664 Blasisaurus canudoi 170 57.17 151 35 2.06 0.34 2.64 0.89 0.66MPZ2010/952 Euhadrosauria indet. 225 65.36 e 112 36 1.68 0.29 1.71 0.50 0.58IPS SRA 27 Pararhabdodon isonensis 243 74.57 104 35 1.44 0.31 1.39 0.43 0.54MPZ2008/258 Arenysaurus ardevoli 275 70.77 95.65 37 1.34 0.26 1.35 0.35 0.65
Notes: French dentaries have not been taken into account, being fragmentary (preserving incomplete dental batteries) or juvenile (with only a small numberof tooth positions). Abbreviations: e, estimated measure;H, height of the dental battery at its midpoint; h, height of coronoid process; L, length of the dentalbattery; n, number of tooth positions; R, relative position of the inflexion point where the ventral margin of the dentary originates (Prieto-Marquez 2010,char. 37); Td, tooth density. Every measurement is given in mm, except for tooth density, which is tooth positions per cm.
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already seen in cranial and postcranial remains, in the
Maastrichtian of the Ibero-Armorican Realm.
Acknowledgements
Financial support was provided by the Ministerio de Economıa yCompetitividad of Spain (projects CGL2010-16447/BTE,CGL2011-30069-C02-01,02/BTE), the Gobierno de Aragon andthe European Social Fund (Grupos Consolidados, Departamentode Educacion y Cultura; JIC, PC-C) and the Gobierno delPrincipado de Asturias (Protocolo CN-04-226; JIR-O). Theexcavations and restoration of the fossils were subsidised by theGobierno de Aragon, the Diputacion Provincial de Huesca and theAyuntamiento de Aren. We thank the editor-in-chief Dr GarethDyke (Ocean and Earth Science, National Oceanography Centreand Institute of Life Sciences, University of Southampton,University of Southampton, Southampton, UK), Dr AndrewMcDonald (Department of Earth and Environmental Science,University of Pennsylvania, Philadelphia, Pennsylvania, USA)and an anonymous reviewer for their comments on themanuscript.Photographs of Figure 5 were taken by Z. Herrera (University ofZaragoza). Rupert Glasgow revised the English grammar.
Notes
1. Present address: CONICET-Instituto de Investigacion enPaleobiologıa y Geologıa, Rıo Negro, Argentina.
2. Email: jigruiz@unizar.es3. Email: rgaeteh@yahoo.es4. Email: violetar@arrakis.es5. Email: joseporiol.oms@uab.es6. Email: jicanudo@unizar.es
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